“Study nature, not books” (Louis Agassiz) Bi 1 Lecture 29 Thursday, June 2, 2005 Evolution 3. Voyages to the Galapagos; The physiology of Diving Mammals 1 Announcements on the Bi 1 Web page: http://www.its.caltech.edu/~bi1/schedule.html Review Session takes place here, today, here, 4 - 6 PM. Sections meet as usual today and tomorrow. The final exam is posted Thursday (today) 6 PM; Due Fri 6/10 4:30PM in the Bi 1 Closet Graduating seniors: papers are due today 5 PM in the Bi 1 Closet 2 Acknowledgements “It takes a village to teach Bi 1 at Caltech” TAs, both grads and undergrads Professional staff: Dr. Jane Mendel, Head TA: Sections and grading Patricia Mindorff, communications David Mathog, Molecular Graphics Eric Slimko, Webmaster Biology Electronics shop computer support: Mike Walsh and Pam Fong Guest lecturers: Robert Freedman David Anderson Cameo appearances from Biology Faculty Advice from other Core Curriculum Resources You, the students 3 The Voyage of the HMS Beagle 1831-1836 Charles Darwin (born 1809), unpaid naturalist Azores Cape Verde Islands Cocos Islands Tahiti Galapagos Islands 35 day visit Canary Isles Bahia Rio de Janeiro Valparaiso Montevideo Cape of Good Hope King George Sound Tasmania Falkland Islands Cape Horn New Zealand journey out journey home 4 Equator Punta Espinoza Age of the archipelago: ~ 1 million yr British Admiralty chart of the Galapagos Islands, based on the Beagle’s observations 5 Darwin’s Finches (5 genera, including Geospizia) • 13 Species, each endemic to the islands • El Nino poses a survival challenge • Highly specialized beaks • Observable evolution in beak size • Distinctive feeding habits cactus finch ground finch tree finch 6 The voyage of the Beagle convinced Darwin that 1. Members of the same species often change slightly in appearance after becoming geographically isolated from each other 2. Organisms living on oceanic islands often resemble organisms found living on a close mainland 3. Factors other than or in addition to climate play a role in the development of plant and animal diversity 4. Organisms of the past and present are related to one another (but there are no fossils in the Galapagos) 7 "The distribution of the tenants of this archipelago would not be nearly so wonderful, if, for instance, one island had a mocking-thrush, and a second island some other quite distinct genus.... But it is the circumstance, that several of the islands possess their own species of tortoise, mockingthrush, finches and numerous plants, these species having the same general habits, occupying analogous situations, and obviously filling the same place in the natural economy of this archipelago, that strikes me with wonder...." Darwin, The Voyage of the Beagle http://www.literature.org/authors/darwin-charles/the-voyage-of-the-beagle/chapter-17.html 8 9 Finch family tree based on a 660-nt sequence Molecular Biology and Evolution 18:299-311 (2001) Darwin’s Galapagos finches .01 change per position 10 Red-Footed Booby (Ben Lester) Blue-Footed Booby 11 Masked booby A recently discovered a behavior, called siblicide, occurs among booby chicks. The larger chick always kills the younger chick, sometimes aided by the parents. Scholars debate the selective advantage of such behavior. Perhaps the parents succeed better by insuring the survival of at least one chick. 12 Male Frigate Bird Displaying no oil on feathers cannot land on water steals other birds’ fish 13 Galapagos Waved Albatrosses in Courtship 2 m wingspan 14 Video of Galapagos Waved Albatrosses in Courtship 15 photo by Ben Lester Female Albatross on the nest 16 photo by Ben Lester 17 The Galapagos tortoises reach sexual maturity at the age of 40 and have clutches of 2 - 26 eggs. Eggs hatch at 85 -180 d. 18 Galapagos Penguins World’s Northernmost population of penguins. After "El Nino" in 1983 the population decreased from 12,000 to just 2,000 birds. 19 Marine Iguanas 20 Marine Iguana feeding Marine iguanas feed once a day. The mature lizards swim out through the tidepools to dive to the bottom for algae; smaller iguanas feed off the rocks in the tidal zone. An iguana may lose up to 10 degrees C of body temperature on these feeding missions. Because they are cold-blooded ("ectothermic") , iguanas must bask on the hot lava rocks throughout the day until they raise their internal temperature. 21 The increased rainfall that accompanies El Niño results in greater food availability for most terrestrial organisms in the Galápagos, but marine life generally suffers from the higher water temperature, which decreases the amount of dissolved oxygen. Green and red algal species, which are the marine iguanas' preferred food, disappear and are replaced in intertidal areas by brown algae which iguanas find hard to digest. Up to 90% of marine iguana populations on islands can die of starvation as a result of these environmental changes. During a recent El Niño event (1997–98), larger individuals of the two island populations shrank more than smaller individuals. The scale of the shrinkage — up to 20% of body length — means that it cannot simply be explained by decreases in cartilage and connective tissue, which together make up only 10% of total body length. Apparently bone absorption accounts for much of the reduction. 22 The fight against salt at Punta Espinoza: marine iguanas cormorants sea lions Marine Iguana 23 salt crystals 24 Flightless Cormorant drying its feathers at Punta Espinoza large flightless birds are common on islands, e.g. kiwi (New Zealand) extinct: Great auk (north Atlantic), dodo (Mauritius), solitaire (Reunion, Rodrigues), moa (New Zealand) 25 California Sea Lions 26 short-eared owl (flies) 27 Flamingos in a salt marsh 28 Pelicans 29 Land iguana (Conolophus sp) 30 Lava lizard doing pushups 31 Territorial iguana chase 32 Sally Lightfoot crab Resembles black crab of Hawaii’s Big Island 33 stingless bee 34 Pinnacle Rock on Bartolome: Excellent diving, lots of marine life 35 The Physiology of Diving 36 Emperor penguins Aptenodytes forsteri 550 m 22 min typical: 2-10 min, 50-500 m 37 Elephant seal Mirounga leonina 1600 m 120 min typical: 20-30 min, 200-800 m 38 Weddell seal Leptonychotes weddellii > 600 m 82 min 39 Bottlenose dolphin Tursiops truncutus 210 m 5 min typical 30 m, 120 s 40 Blue whale Balaenoptera musculus 300 m 50 min 41 Sperm whale Physeter macrocephalus 3000 m 90 min 42 In metabolism, electrons move around inside cells until they reach oxygen, which has the highest electron affinity of the biological elements Little Alberts 2-7 © Garland 43 Diving mammals must store oxygen in order to conduct aerobic metabolism Emperor penguin Baikal seal Weddell seal In the absence of oxygen, glucose pyruvate and lactic acid. ~ 2 moles of ADP are converted to ATP. In the presence of oxygen, glucose + O2 H20 + CO2. ~ 24 moles of ADP are converted to ATP enzymes and carriers transfer 40% of the bond energy to high-energy phosphate bonds, in small steps. 44 Diving mammals have much higher: blood volume hemoglobin concentration and especially myoglobin two oxygen-carrying heme proteins than other mammals. More than 80% of the oxygen is in the blood and muscle. 45 Diving mammals have much higher: blood volume hemoglobin concentration and especially myoglobin two oxygen-carrying heme proteins than other mammals. More than 80% of the oxygen is in the blood and muscle. 46 Crittercam (UC Santa Cruz) 35 cm detachable CCD lens camera 900 nm LEDs batteries and 8 mm tape recorder computer float transmitter Ti or Al housing fins Audio channels (1) accelerometer (2) microphone Transducers for pressure, water speed, and compass bearing are sampled once per second, and the data are stored on a PCMCIA card. A separate housing (17 cm long and 5.5 cm in diameter) for the gimbaled flux-gate compass is positioned behind the main housing and connected to it with a cable. 47 Diving mammals glide much of the time 48 Diving mammals save ~ 28% of their energy by gliding (plexiglas dome) 49 Bottlenose dolphins save energy by slowing their heart rate during a dive 50 Pressure effects on air gases 10 m = 1 bar Sport diving is limited to ~ 40 m 1. The bends: N2 bubbles form in the blood 2. Nitrogen narcosis 3. Oxygen toxicity 51 Tanks 200 bar Regulators 52 Diving mammals don’t get the bends: adaptations to pressure “Lungs are a liability for deep divers because, in contrast to muscle and blood, they are a better nitrogen store than oxygen store.” a. flexible chest; b. no collagen in the smallest branches “The early occurrence of lung collapse in seals makes the lung almost useless as an O2 store, whereas it limits N2 absorption during the dive.” Some compressed air remains in the windpipes, but a collagen lining prevents absorption by the blood complete peripheral collapse at 20-50 m 53 Other effects of water’s high density 1. Heat capacity and heat conduction Diving mammals solve this problem with blubber. Scuba divers use exposure suits (wet suit, dry suit). Both of these materials compress at depth, increasing the animal’s density and providing negative buoyancy. 2. Greater index of refraction 3. Greater speed of sound 4. Greater light absorption 54 2. Objects look larger to a diver (Phys 2a) apparent object real object q’ q’ water n’ ~ 1.3 q air in mask n=1 q nsinq = n’sinq’ 55 3. A diver cannot judge sound direction delay between the two ears ~ 7 sin q ms (can be determined to within ~ 10 ms) q Sound travels 3-4 times faster in water 20 cm 0.2 m/(300 m/s) ~ 7 ms 56 4. Objects look blue to a diver a few m of water absorbs long wavelengths shorter wavelengths penetrate further 57 A Summary of Adaptations in Diving Mammals 1. Compressible lungs: decrease buoyancy, decrease bends 2. High myoglobin concentration 3. Blubber 4. “Intermittent locomotion” = gliding 6. Decreased heart rate decreases heat flow 5. Wave riding and jumping 6. Countercurrent cooling devices for testes 58 Darwin, The Voyage of the Beagle Darwin, The Origin of Species Michael Jackson, Galapagos, University of Calgary Press, 1994 Jonathan Weiner, The Beak of the Finch: a Story of Evolution in our Time, Knopf, 1995 59